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North American Fungi Volume 5, Number 5, Pages 9-21 Published December 22, 2010 Amanita in the Rocky Mountain alpine zone, USA: New records for A. nivalis and A. groenlandica Cathy L. Cripps1 and Egon Horak2 1Plant Sciences and Plant Pathology Department, Montana State University, Bozeman, MT 59717-3150, U.S.A; 2Nikodemweg 5, AT-6020 Innsbruck, Austria Cripps, C. L., and E. Horak. 2010. Amanita in the Rocky Mountain alpine zone, USA:New records for A. nivalis and A. groenlandica. North American Fungi 5(4): 9-21. doi:10.2509/naf2010.005.0052 Corresponding author: Cathy L. Cripps, [email protected]. Accepted for publication June 20, 2010. http://pnwfungi.org Copyright © 2010 Pacific Northwest Fungi Project. All rights reserved. Abstract: A limited number of Amanita species have been reported from cold dominated arctic-alpine environments, primarily with dwarf and shrub willows. This includes reports from Alaska, Canada, Greenland, Iceland, Scandinavia, Scotland, the Alps, and Russia. Here we report Amanita nivalis, A. groenlandica f. alpina n. f. and A. vaginata above tree line in the Rocky Mountains with Salix reticulata, S. nivalis, S. glauca, and S. planifolia. The distribution of Amanita groenlandica is extended to the north-central Rockies, and more definitively to the Beartooth Plateau at 3,100 to 3,400 m a.s.l. A new alpine form is described for this arctic species. Amanita nivalis is confirmed from numerous cirques and high passes 3,600-3,900 m a.s.l. in the southern Rocky Mountains (Front Range, Sawatch Range, San Juan Mountains), adding a disjunct component to its circumpolar distribution. These localities are thousands of miles from the Arctic and are likely near the southern-most extent of its distribution in North America. Rocky Mountain species are reported in context with arctic-alpine Amanita species from the Northern Hemisphere. Key words: Agaricales, Amanitaceae, Amanita groenlandica, Amanita nivalis, alpine, Arctic, cold- climate fungi, ectomycorrhizal, Rocky Mountains, Salix, tree line 10 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 Introduction: The genus Amanita is known from the krummholz conifer zone. All are from arctic-alpine habitats throughout the ectomycorrhizal species associated with alpine Northern Hemisphere (Favre 1955, Gardes and vegetation including shrub willows Salix glauca Dahlberg 1996, Gulden et al. 1985, Hallgrimsson L. and S. planifolia Pursh., and dwarf willows S. 1979, Hutchison et al. 1988, Knudsen & Borgen reticulata L. and S. arctica Pall. Three species of 1987, Miller et al. 1982, Watling 1985, 1987), Amanita are reported here from the Rocky however, there have been no formal reports of Mountain alpine zone along with their ecology strictly alpine species from the Rocky Mountains. and in context with other arctic-alpine taxa in Only a limited number of Amanita species occur this genus. in cold dominated environments mainly in association with dwarf/shrub willows and dwarf birch. At least one species (A. nivalis) has a Materials and Methods northern circumpolar distribution pattern. Study Area Historical taxonomic confusion and unresolved The Rocky Mountains extend along the spine of synonymies have made it unclear as to the exact North America continuing 5,000 km north from number of distinct arctic and alpine taxa of New Mexico through Canada to Alaska. The Amanita. To add to the confusion, the Rocky Mountain alpine tundra, although basidiomes of many alpine Amanita species are fragmented on mountain tops, constitutes a white, near white, or possess a whitish pileus phytogeographically significant region, under some environmental conditions. Amanitas comprised of the Northern, Middle, and Southern often occur as single basidiomes in alpine floristic provinces across a range of latitudes habitats so intraspecific variation cannot be (Arno and Hammerly 1984). Our study examined easily examined. Harsh arctic and alpine sites south of the Canadian border, above treeline conditions can quickly destroy salient features, at 3,000 m a.s.l. in the north at latitude 45°N which is problematic in macromorphological (Montana and Wyoming) and above treeline at evaluation. A majority of arctic-alpine Amanita 3,700 m a.s.l. in southern Colorado at latitudes of species are in sect. Vaginata characterized by 37°-39°N. Portions of the alpine areas in both of basidiomes with a saccate volva, absence of an these regions escaped glaciation. annulus and non-amyloid spores (Tulloss 1994). Species within this group often lack distinct Middle-Northern Floristic zone: The Beartooth features that allow clear taxonomic separation, Plateau is our primary study area for this floristic although molecular work has helped define zone, and extends from Montana into northern subalpine species (Drehmel et al. 1999, Moncalvo Wyoming. The plateau is one of 11 situated along et al. 2000, Weiss et al. 1998). the MT-WY border that together comprise the most extensive alpine tundra in the lower 48 An extensive NSF-sponsored survey of alpine states (Anderson 1994). The Beartooth Plateau fungi along the Rocky Mountain cordillera for the itself is about 20 km long by 10 km wide, at lat. Beartooth Plateau on the Montana/Wyoming 450N and long 109°W. The vast sweeping gentle border and the southern floristic zone in slopes are covered by an intricate patchwork of Colorado resulted in over 2,000 collections of meadows, shrubby wetlands, grasslands and fell- alpine agarics that fall into ca. 200 species fields (Cooper et al. 1997, Eversman 1995, designations (Cripps and Horak 2006a, 2006b, Johnson and Billings 1962). Details of the 2008, Osmundson et al. 2005). Amanita species vegetation and soils are in Anderson (1994), were regularly collected as part of this survey Bamberg and Major (1968), and Scott (1995). from habitats above treeline and strictly away Soils are primarily derived from granitic material. Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 11 The collecting season for fungi is typically from drawn, and identified on subsequent completion mid-July to the end of August. of microscopic examination. Spores and other microscopic features were examined in Southern Floristic Zone: The southern Rocky 3% KOH after rehydration. Colors are from Mountains are isolated from the northern ranges Kornerup and Wanscher (1967). Voucher by the Wyoming Plateau. The individual study specimens are in herbaria at Montana State sites lie in a chain along the Continental Divide at University, Bozeman, MT (MONT) and the elevations from 3600 to 4000 m a.s.l. at lat. 36°- Institute of Integrative Biology, ETH, Zurich, 38°N. The sites include high mountain passes, Switzerland (ZT). An advantage for our study was unique cirques, and basins with a wealth of that particular environmental conditions (high habitats including open windswept slopes, precipitation at local fungus peak season) often shrubby wetlands, extensive dwarf Salix and resulted in large fruitings that provided an Dryas patches, meadows, turf, and fell-fields. A opportunity to examine basidiome variation variety of soil types occur in the mountain ranges within the main alpine taxa that occur in the from granitic to limestone. The fruiting season is Rockies. from mid-July through August or into September after which snow can close the high areas. Results: Numerous basidiomes of Amanita were observed from 1999 and 2008 on the Ectomycorrhizal Vascular Plants: The primary Beartooth Plateau sites (MT/WY) above 3,000 m ectomycorrhizal plants of these alpine areas a.s.l. (Fig. 1) and on the southern sites (Colorado) include dwarf willow species S. reticulata, S. above 3,700 m a.s.l. The records represent three arctica and more rarely S. rotundifolia Trautv. species of Amanita that occur above tree line (not and S. cascadensis Cockll. (Cripps and Eddington in association with conifers) in the Rocky 2005, Scott 1995). Shrub willows consist Mountains. Only North American collections are primarily of S. glauca which is more common in described here with emphasis on local ecology southern areas and S. planifolia which is more and distribution patterns (Table 1). prevalent in the north. Betula glandulosa Michx. (= B. nana L.) is rare in the Rocky Mountain alpine zone and no Amanita species were recorded with this host. Dryas octopetala L. mats can be extensive in the Rocky Mountain alpine belt and host a limited diversity of ectomycorrhizal fungi (Cripps and Horak 2006b), however, to date no Amanita species have been recorded in strict association with Dryas for our sites. Collection and processing Basidiomes were collected from 1999 to 2008, primarily on the Beartooth Plateau in the North- Central Rockies and on the Front Range, Sawatch Range and San Juan Mountains to the South. Each collection was photographed, described, Fig. 1. Alpine area of Beartooth Plateau, Rocky Mountains, MT/WY, USA. 12 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 Table 1. Collection data for Amanita species in the Rocky Mountain alpine zone: A. groenlandica collections are from the Beartooth Plateau* near the Montana and Wyoming border, A. nivalis collections from Colorado**, and A. vaginata from the Beartooths. Cripps (=CLC) collections are in MONT and Horak (=ZT) collections are in ETHZ. Collection Date Range Location Elev. Hosts Amanita groenlandica f. alpina Horak 6410 07 Aug-98 Beartooth Site 1: Birch site 3,100 Salix reticulata Cripps 1213 08 Aug-98 Beartooth Site 2: Highline trail 3,100 Salix spp. Cripps 1377 19 Aug-99 Beartooth Site 1: Birch site 3,000 S. reticulata & S. planifolia Cripps 1392 20 Aug-99 Beartooth Site 2: Highline trail 3,100 Salix reticulata Cripps 1405 21 Aug-99 Beartooth Site 3: Frozen lakes 3,100 Salix glauca/planifolia Cripps 1413 18 July-00 Beartooth Site 2: Highline trail 3,100 Salix reticulata Cripps 1585 20 July-01 Beartooth Site 2: Highline trail 3,100 Salix reticulata Cripps 1779 21 Aug-02 Beartooth Site 4: Headwall 3,400 Salix reticulata Cripps 1929 28 Aug-02 Beartooth Site 2: Highline trail 3,100 Salix cf planifolia Cripps 1970 12 Aug-03 Beartooth Site 5: Top of Pass 3,400 Salix reticulata & S.